Water and Nitrogen Uptake in Upland and Swamp Rice

The water uptake per unit dry matter and nitrogen uptake perunit water absorbed of swamp rice (Dima) and upland rice (Kindinga)at three nitrogen levels equivalent to 23.6, 70.8, and 118,kg./ha. (20, 60 and 100 lb./acre) respectively were studiedunder tropical green-house conditions in Trinidad. The water uptake per unit dry matter produced was significantlygreater in Kindinga than in Dima at the three nitrogen levels.Additional nitrogen increased the water requirement of Kindingabut decreased that of Dima, the greater increase in Kindingabeing attributed to increase in leaf area. The efficiency of nitrogen absorption in terms of water transpiredwas greater in Dima than in Kindinga at the three nitrogen levels.     

The Contribution of Different Organs to Grain Weight in Upland and Swamp Rice

The contribution of dry matter was significantly greater inthe ears of Dima than that of Kindinga. The flag-leaf lamina,flag-leaf sheath and peduncle, lower leaves and stem of Kindingamade greater contribution of dry matter in the grain at harvestthan those of Dima. On the average of the two varieties 23 percent. of the dry matter in the grain at harvest originated fromphotosynthesis in the ears; 60 per cent. came from photosynthesisin the flag-leaf lamina and sheath and peduncle and 17 per cent.from parts of the shoot below the flag leaf.     

氮肥处理对氮素高效吸收水稻根系性状及氮肥利用率的影响

2011—2012年在土培条件下,以氮素吸收效率差异较大的15个常规籼稻为供试材料,研究氮肥运筹对不同氮效率品种根系性状、成熟期吸氮量及氮肥利用率的影响,分析影响氮高效水稻氮素吸收的主要根系性状。结果表明:(1)各氮肥处理下,成熟期吸氮量均表现为氮高效品种氮中效品种氮低效品种。适量增施氮肥及基肥+促花肥处理有利于氮高效品种吸氮量的增加,氮素吸收受品种、氮肥处理的显著影响。(2)在施氮量处理下,氮高效品种单株不定根数、单株根干重、单株不定根总长大或较大,单株根活力在常氮(N2)、高氮(N3)处理下有一定的优势;在施氮时期处理下,氮高效品种单株不定根数、单株不定根总长、单株根干重、单株根系总吸收面积、单株根系活跃吸收面积、抽穗期冠根比多数处理有优势;增施氮肥有利于促进氮高效品种单株不定根总长和单株根活力的提高,适量施氮有利于单株不定根数、单株根干重增加,前期施氮可促进不定根的发生和伸长,后期施氮有利于不定根的充实和根系生理性状的提高。此外,增施氮肥可提高各类品种冠根比;(3)在常氮、高氮处理下,氮高效品种氮肥利用率大于氮中效、氮低效品种。(4)提高单株不定根数、单株不定根总长、单株根活力及抽穗期冠根比有利于各类品种吸氮量的提高,增加根干重对氮高效品种吸氮量的提高也有显著的促进作用。结合相关分析与通径分析结果,抽穗期冠根比及单株不定根数、单株根活力、单株不定根总长、单株根干重是影响氮高效品种吸氮能力的主要根系性状。     

Effects of growth regulators, CCC and B9, on some potato varieties

The effects of CCC and B 9 on the growth habit of potato differed between varieties. CCC diminished stem lengths and dry weight more than Bo because CCC was applied early when shoots emerged from the soil, but B 9 was applied about 3 weeks later when several leaves had formed. In some varieties lateral stem growth was increased by treatment and in others decreased. There was an inverse relation between main stem and lateral stem growth so that varieties with vigorous main stem growth had poor lateral growth and vice-versa. Treatment with the growth-regulators diminished leaf dry weight of main stem leaves less than leaf area, but the degree of magnitudes of the changes depended on the variety. Both regulators lessened net assimilation rate. Net assimilation rate and dry matter per unit area of leaf were inversely related, possibly because accumulation of substances in leaves decreases photosynthesis. Stolon dry weight was positively correlated with main stem leaf area. There was a direct relation between stem length and tuber dry weight, suggesting that tuber initiation occurs at different stem lengths in different varieties. Tubering was earliest in Epicure and latest in King Edward. Epicure had the greatest tuber weight and smallest stem length.     

Canopy development of a determinate and an indeterminate cultivar of Vicia faba L. under contrasting plant distributions and densities

Field experiments were conducted in 1987 and 1988 to quantify differences in canopy formation between an indeterminate and a determinate genotype of Vicia faba L., grown at two plant densities and three spatial distributions. The number of stems per unit area produced by determinate plants was related to the growth rate before flowering. Leaf production per stem per unit of thermal time was similar in both plant types, but twice as many leaves per stem were produced by the indeterminate cultivar. The indeterminate cultivar produced fewer and smaller leaves in the warmer and drier weather of 1988 than in 1987. The determinate genotype produced similar sizes and numbers of leaves in both years, but fewer tillers developed in 1988 than in 1987. Accordingly, leaf mass per unit ground area was greater in 1987 than in 1988 in both genotypes. Except during early flowering, relationships between leaf mass and leaf area were constant, with higher specific leaf areas in the determinate than the indeterminate genotype. Shoot dry matter partitioning into leaves was identical in both years for indeterminate plants, but differed in determinate ones.
It is concluded that canopy development is regulated through individual leaf weight and leaf number per stem in non-tillering indeterminate, and by stem numbers per unit area in tillering determinate plants.     

The relation between some growth parameters and yield components of rice as influenced by nitrogen level and genotype

The effects of three levels of nitrogen (0, 2 and 4 g (NH₄)₂SO₄ per pot) were evaluated in four rice varieties, differing in grain size, under flooded conditions in a tropical environment. Addition of nitrogen increased plant height and leaf number on the main culm, and hastened flowering significantly in all varieties. It also increased the dry weights of leaf, and culm plus sheath at the maximum tillering stage, anthesis and harvest. The average dry weights of both leaf, and culm plus sheath increased significantly in all varieties until anthesis. Following anthesis, the dry weights of both leaf, and culm plus sheath of the large-seeded varieties, Zhu Lian and PP₆R, further increased mainly due to emergence of the late tillers, but decreased in the small-seeded varieties, IR 747 and P₃C₂₆. During grain-filling, vars Zhu Lian and PP₆R accumulated dry matter more than the other two varieties and did not use their reserve starch present in the culm plus sheath, presumably because of the lack of demand. By contrast, current photosynthesis and ‘reserve’ contributed about 70% and 30%, respectively, of the grain yield in IR 747 and P₃C₂₆. Addition of nitrogen increased grain yield significantly in all varieties. The yield increase in IR 747 and PP₆R was essentially due to increase in grain number, but that in P₃C₂₆ and Zhu Lian was due to increases in both grain size and grain number, the latter having much greater influence. Neither the percentage of filled spikelets nor the 1000-grain weight was influenced by nitrogen addition. It was concluded that grain number is the single most important yield component in rice. Path-coefficient analysis indicated that in a breeding programme involving these four rice varieties, the cross between Zhu Lian and P₃C₂₆ would be most successful in an effort to combine high spikelet number and large grain size.     

Heteroses in Net Photosynthetic Rate, Leaf Area, Tillering, and Some Physiological Characters of 35 F1 Rice Hybrids

Net photosynthetic rate, leaf area, tillering, shoot weight,and leaf area ratio for 35 F₁ rice (Oryza sativa L.) hybridsduring the vegetative stage were compared with correspondingvalues for parental varieties. Shoot weight and leaf area showedheterosis. However, hybrid vigour was not reflected in net photosyntheticrate. Leaf area was closely correlated with heteroses in shootdry weight and tillering. Thus F₁ rice hybrids grow vigorouslybecause of their high leaf area development, which is causedby high tillering. Key words: F₁ rice hybrid, Net photosynthetic rate, Leaf area     

Nitrogen supply and the control of expansive growth and function in leaves and roots

In plants which have acclimatized to limiting supplies of nitrogen (steady-state nutrition), leaf expansion (numbers and sizes of leaves and cells) is under tight control. Over a wide range of nitrogen supplies, the control of leaf growth is associated with a narrow band of photosynthetic rate per leaf area (measured at the growth climate) and, at limiting supplies, a carbon uptake which is in excess of immediate carbon usage in structural growth.For every increment of nitrogen absorbed, root extension is greater at limited nitrate supply, but V_max values (per root dry weight) for nitrate absorption are typically less. However, the capacity of the whole root system for nitrate uptake at limited supply is sufficient to allow for maximum growth, should nitrate supply be increased.It is concluded that a better understanding at the cellular level of the mechanisms which result in a greater inhibition of the expansion of single leaves than of root extension would contribute to an understanding of differences in carbon sink strength among plant organs. This may be a crucial step towards a more physiologically-based appreciation of plant dry matter distribution among organs in plants experiencing different nitrogen supplies.     

Carbon and nitrogen economy of 24 wild species differing in relative growth rate

The relation between interspecific variation in relative growth rate and carbon and nitrogen economy was investigated. Twentyfour wild species were grown in a growth chamber with a nonlimiting nutrient supply and growth, whole plant photosynthesis, shoot respiration, and root respiration were determined. No correlation was found between the relative growth rate of these species and their rate of photosynthesis expressed on a leaf area basis. There was a positive correlation, however, with the rate of photosynthesis expressed per unit leaf dry weight. Also the rates of shoot and root respiration per unit dry weight correlated positively with relative growth rate. Due to a higher ratio between leaf area and plant weight (leaf area ratio) fast growing species were able to fix relatively more carbon per unit plant weight and used proportionally less of the total amount of assimilates in respiration. Fast growing species had a higher total organic nitrogen concentration per unit plant weight, allocated more nitrogen to the leaves and had a higher photosynthetic nitrogen-use efficiency, i.e. a higher rate of photosynthesis per unit organic nitrogen in the leaves. Consequently, their nitrogen productivity, the growth rate per unit organic nitrogen in the plant and per day, was higher compared with that of slow growing species.     

烟草不同基因型耐低氮能力差异评价

以单株叶重及其氮素反应指数作为耐低氮能力的评价指标,分析了烟草种质资源的耐低氮能力以及单株叶重及其氮素反应指数与主要农艺性状间的相关关系。结果表明,不同施氮水平下多数农艺性状基因型间差异较大,在低氮水平下表现最大;和其他农艺性状相比,单株叶重在不同氮素水平间平均差异最大。不同施氮水平下,单株叶重均与最大叶叶面积、单株生物量呈显著或极显著正相关;单株叶重的氮素反应指数与单株叶重、生物量呈显著或极显著正相关,且低氮水平下单株叶重的氮素反应指数与上述性状间的相关性比中氮水平下更为密切。在供试的36个烟草品种中,永定400号、金烟6号、红花大金元、G80、Nc82等烟草种质具有较强的耐低氮能力,可以作为氮高效的烟草品种利用。     

Physiological Causes of Differences in Grain Yield between Varieties of Barley

In a field experiment on barley at Rothamsted with the highmean yield of 49 cwt. of grain per acre, the varieties Proctorand Herta produced 10—15 per cent, more grain than Plumage-Archeron plots that received no nitrogenous fertilizer. When nitrogenwas applied the difference was increased to about 30 per cent.,because the higher nitrogen supply caused the Plumage-Archercrop to lodge and did not increase its yield, while Proctorand Herta remained standing. The three varieties did not differ in leaf-area index nor innet assimilation rate before ear emergence, so that all hadthe same total dry weight. After ear emergence, the leaf-areaindices of Proctor and Plumage-Archer were nearly equal, butthat of Herta was smaller. Assuming that the photosyntheticefficiency of the leaves continued to be the same in all varieties,the higher grain yields of Proctor and Herta cannot be attributedto greater production of dry matter by the leaves, either beforeor after ear emergence. A pot experiment on plants with shadedears confirmed that the dry matter contributed to grain yieldby unit leaf area was nearly equal in all the varieties. The higher grain yield of Proctor and Herta than of Plumage-Archermust therefore have come from additional photosynthesis in partsof the plant other than the leaves, i.e. in the ears themselves.An attempt to demonstrate this directly in a pot experiment,by comparing the grain yields of plants with shaded or withunshaded ears, was unsuccessful because the varieties behaveddifferently in pots; Proctor and Herta produced only about 6per cent, more grain yield than Plumage-Archer, and though thedecrease in grain yield by shading the ears was slightly greaterfor Proctor and Herta, the differences were not significant. The sum of ear sizes (estimated from length and breadth measurements)per m.1 in the field experiment was greater for Proctor andHerta than for Plumage-Archer. Also the distribution of drymatter between developing ears and shoots apparently differedwith variety, so that at ear emergence the dry weight of earsper m.² was greater in the two higher yielding varieties. Theincreased amount of photosynthetic tissue in the ears of Proctorand Herta, as measured by size or weight, may not wholly explaintheir greater dry-matter production; ears of Herta may alsohave a higher photosynthetic efficiency. No differences in nutrient uptake that could account for thevarietal differences in grain yield were found. Plumage-Archerabsorbed more potassium, and Herta less phosphorus than theother varieties. About a quarter of the final content of nitrogen,and a third of the phosphorus, was absorbed after ear emergence,but the potassium content was nearly maximal at ear emergenceand later decreased. The pot experiment showed that, on the average of all varieties,26 per cent. of the dry matter in the grain at harvest originatedfrom photosynthesis in the ears, including 10 per cent, fromthe awns; 59 per cent, came from photosynthesis in the flag-leaflamina and sheath and peduncle, and 15 per cent, from partsof the shoot below the flag leaf.     

Leaf Lateral Asymmetry in Morphological and Physiological Traits of Rice Plant

Leaf lateral asymmetry in width and thickness has been reported previously in rice. However, the differences between the wide and narrow sides of leaf blade in other leaf morphological and physiological traits were not known. This study was conducted to quantify leaf lateral asymmetry in leaf width, leaf thickness, specific leaf weight (SLW), leaf nitrogen (N) concentration based on dry weight (Nw) and leaf area (Na), and chlorophyll meter reading (SPAD). Leaf morphological and physiological traits of the two lateral halves of the top three leaves at heading stage were measured on 23 rice varieties grown in three growing seasons in two locations. Leaf lateral asymmetry was observed in leaf width, leaf thickness, Nw, Na, and SPAD, but not in SLW. On average, the leaf width of the wide side was about 17% higher than that of the narrow side. The wide side had higher leaf thickness than the narrow side whereas the narrow side had higher Nw, Na, and SPAD than the wide side. We conclude that the narrow side of leaf blade maintained higher leaf N status than the wide side based on all N-related parameters, which implies a possibility of leaf lateral asymmetry in photosynthetic rate in rice plant.     

Effect of nitrogen supply on growth,allocation and gas exchange characteristics of two perennial grasses from inland dunes

Summary We studied the effects of nitrogen supply on growth, allocation, and gas exchange characteristics of two perennial grasses of dry, nutrient-poor inland dunes: Corynephorus canescens (L.) Beauv. and Agrostis vinealis Schreber. C. canescens invests more biomass in leaves and less in roots, but has less leaf area and more root length per unit plant weight than A. vinealis. A. vinealis invests more nitrogen per unit leaf weight, but less per unit leaf area, despite a similar relative nitrogen investment in leaves and plant nitrogen concentration. Between-species differences in the rate of net photosynthesis, transpiration and shoot respiration are positively related to leaf nitrogen content per unit leaf area. The rate of net photosynthesis per unit plant weight is higher for A. vinealis at both levels of nitrogen supply, due to differences in leaf area ratio (LAR), and despite the reverse differences in the rate of net photosynthesis per unit leaf area. The water use efficiency of the two species is similar and increases significantly with an increase in nitrogen supply. The photosynthetic nitrogen use efficiency on the other hand is not affected by nitrogen supply, while at both low and high nitrogen supply A. vinealis has a 10% higher photosynthetic nitrogen use efficiency than C. canescens.     

红豆杉幼树异龄叶的功能性状

植物叶片功能性状随叶龄的变化体现了植物的生活史策略,反映着植物对有限资源的权衡与配置特征.本研究以红豆杉幼树异龄叶为对象,测定了3个年龄段叶片的14个功能性状,分析红豆杉叶片老化过程中生态策略的调整方式.结果 表明:1年生叶的比叶面积显著大于2、3年生叶,3年生叶的叶厚、叶面积、叶体积、叶鲜重和叶干重显著大于1、2年生...     

Interactions of Nitrogen, Phosphorus, and Potassium supplied in Leaf Sprays or in Fertilizer added to the Soil

When sugar-beet plants grown in pots were sprayed daily withnutrient solutions supplying nitrogen, phosphorus, and potassiumseparately or in all combinations, with precautions to preventspray falling on the soil in which the plants were grown, allthree nutrients were absorbed through the leaves. In one experimentnitrogen and potassium, and in another only nitrogen, causedincreases in plant dry weight and leaf area. Swedes absorbedphosphorus from leaf sprays and from fertilizer applied to thesoil, but only the fertilizer caused an increase in dry weight. Absorption of any of the nutrients tested from a spray containingmore than one nutrient was unaffected by the presence of othersin the spray, but spraying with nitrogen-containing solutionsincreased the absorption of phosphorus and potassium from thesoil, and potassium in sprays increased the uptake of phosphorusfrom the soil. Nitrogenous fertilizer applied to the soil increased the leafarea of sugar-beet plants, and hence it also increased the amountsof nitrogen, phosphorus, and potassium deposited on the leaveswhen they were sprayed with solutions of these nutrients, andthe amounts absorbed from the spray into the plants. Phosphaticfertilizer had no effect on uptake from leaf sprays. Potassicfertilizer did not affect leaf area or the estimated volumeof spray solution retained on the leaves, but it appeared toreduce uptake of potassium from the spray. Dry weight per plant was increased by all three nutrients infertilizer, and sugar yield of the roots was increased by nitrogenand potassium in fertilizer, and by nitrogen in spray. Applicationof a nutrient in leaf spray reduced the responses in dry weightand sugar yield to the same nutrient applied in fertilizer tothe soil. Less nitrogen, but more phosphorus, was taken up from the leafsprays than from fertilizer. Nutrients from sprays producedsmaller increases in total dry weight and in dry weight perunit of absorbed nutrient than the same nutrient from fertilizer. The apparent percentage recovery of nitrogen applied in spray,based on estimates of the volumes of solution retained on theleaves, was unaffected by fertilizer treatment, that of phosphoruswas increased by nitrogen fertilizer, and that of potassiumwas increased by nitrogen fertilizer and reduced by potassiumfertilizer. The volume of spray solution held on the leaveswas probably overestimated, so that the highest apparent recovery,about 60 per cent., may represent an almost complete true recovery,because only trivial amounts of the nutrients that had beenapplied in spray remained on the leaf surface to be removedby washing before harvest. Lower apparent recoveries may bedue to reduced uptake from the soil of the nutrient suppliedin spray.     

Hydraulic conductance as well as nitrogen accumulation plays a role in the higher rate of leaf photosynthesis of the most productive variety of rice in Japan

An indica variety Takanari is known as one of the most productive rice varieties in Japan and consistently produces 20-30% heavier dry matter during ripening than Japanese commercial varieties in the field. The higher rate of photosynthesis of individual leaves during ripening has been recognized in Takanari. By using pot-grown plants under conditions of minimal mutual shading, it was confirmed that the higher rate of leaf photosynthesis is responsible for the higher dry matter production after heading in Takanari as compared with a japonica variety, Koshihikari. The rate of leaf photosynthesis and shoot dry weight became larger in Takanari after the panicle formation and heading stages, respectively, than in Koshihikari. Roots grew rapidly in the panicle formation stage until heading in Takanari compared with Koshihikari. The higher rate of leaf photosynthesis in Takanari resulted not only from the higher content of leaf nitrogen, which was caused by its elevated capacity for nitrogen accumulation, but also from higher stomatal conductance. When measured under light-saturated conditions, stomatal conductance was already decreased due to the reduction in leaf water potential in Koshihikari even under conditions of a relatively small difference in leaf-air vapour pressure difference. In contrast, the higher stomatal conductance was supported by the maintenance of higher leaf water potential through the higher hydraulic conductance in Takanari with the larger area of root surface. However, no increase in root hydraulic conductivity was expected in Takanari. The larger root surface area of Takanari might be a target trait in future rice breeding for increasing dry matter production.     

Effects of Night-break Treatment on Leaf Area and Leaf Dry Weight in Callistephus chinensis

Plants of Callistephus chinensis were grown in eight hours ofdaylight per day and received either uninterrupted dark periodsor dark periods interrupted by one hour of low-intensity light.The growth in area and the accumulation of dry matter was followedfor leaves at different heights of insertion on the stem. Lightinterruption treatment caused more rapid surface expansion ofindividual leaves. The rate of accumulation of dry matter wasalso increased but less in proportion to the increase in areaso that the area per unit weight of leaf material, i.e. thespecific leaf area, was greater. The final areas and dry weightsof leaves were also considerably greater than in the uninterruptednight treatment. When plants were transferred from uninterruptedto interrupted nights, a similar response was obtained fromthose leaves which began to expand rapidly after the commencementof the treatment. The reciprocal transfer into uninterrupted nights immediatelyretarded the rate of leaf expansion and final leaf areas wereless. The rate of accumulation of dry matter was not affectedso rapidly and the area per unit weight of leaf material fellbelow that recorded from leaves which had always received uninterruptednights. Within any one treatment and on any one occasion thespecific leaf area was almost constant, regardless of heightof insertion and therefore also of leaf size and degree of development. It is postulated that a promotor of leaf growth is producedin the light interruption treatment and that this regulatormay increase cell expansion. Transfer to uninterrupted nightsmay halt the synthesis of this regulator and may also lead tothe production of an inhibitor of leaf expansion. Finally, it is concluded from a survey of the literature, that,in general, where a response to photoperiod exists, leaf areais greater in long days than in short days.     

褐飞虱侵害后不同水稻品种根及叶片脱落酸含量的变化

为了解褐飞虱Nilaparvata lugens （Stål）侵害后水稻耐虫性与植物体内源激素关系,应用酶联免疫吸附法（enzyme-linked immunosorbent assay, ELISA）研究褐飞虱若虫侵害分蘖期超级培矮64S/E32和TN1,灌浆期协优963和TN1后根及叶片脱落酸（abscisic acid, ABA）含量变化.结果表明：褐飞虱侵害分蘖期超级培矮64S/E32和TN1后3 d,叶片ABA含量显著上升,ABA含量根冠比（根ABA/叶片ABA）显著下降;侵害后6 d,超级培矮64S/E32叶片ABA含量显著下降,根冠比显著上升;但TN1叶片ABA含量在褐飞虱侵害后3 d和6 d显著上升,根冠比显著下降.褐飞虱侵害灌浆期协优963与分蘖期超级培矮64S/E32变化一致,TN1在褐飞虱侵害后3 d叶片ABA含量显著上升,根冠比显著下降;侵害后6 d,叶片ABA含量、ABA含量根冠比均显著上升.由ABA含量变化百分比可见,分蘖期ABA含量变化幅度较灌浆期大;耐虫品种变化幅度较感虫品种大,持续期较感虫品种短;叶片变化幅度较根部大.褐飞虱侵害后,两种不同生育期两种抗性不同的水稻品种比较,耐虫品种叶片ABA含量先上升（3 d）后下降（6 d）,ABA含量根冠比先下降（3 d）后上升（6 d）;感虫品种叶片ABA含量持续上升（3 d和6 d）,分蘖期ABA含量根冠比持续下降（3 d和6 d）,灌浆期ABA含量根冠比先下降（3 d）后上升（6 d）;耐、感虫水稻品种根部变化规律不明显.这些差别表明不同水稻（耐虫和感虫）品种受褐飞虱侵害后体内ABA含量变化规律不同.本研究结果对深入阐明水稻耐虫品种的机制具有重要参考价值.     

Effect of Defoliation on the Growth and Yield of Wheat

The effect of removal of the fifth (L₁), fifth and sixth (L₂) and fifth, sixth and seventh leaves (L₃) from the main shoot and of similar number of leaves from the primary and secondary tillers, on the growth and yield of two varieties of wheat, was investigated. The plants were grown in pots under natural conditions. Defoliation did not affect tillering, ear number, and number of spikelets per ear. The area of the last 3 leaves was reduced only in L₃. Defoliation reduced the growth of stem up to the flag leaf stage but thereafter there was a remarkable recovery in increase in height and dry weight. The grain yield of the main shoot was affected relatively more than that of the primary and secondary tillers by defoliation. The grain yield per plant was reduced by 5 per cent in L₁, and L₂ and by 11 per cent in L₃. The bearing of these observations on productivity per unit area is indicated and it is suggested that a plant with two fewer leaves per shoot may suffer less from mutual shading of leaves and hence prove more efficient in a dense community.     

Competition between grass and clover in spring as affected by nitrogen fertiliser

Detailed measurements of irrigated ryegrass/white clover swards growing without interruption, with or without nitrogen fertiliser in spring, showed that the relative growth rate of clover was as great as that of grass, in the + N sward, and considerably greater than grass, in the – N sward. Clover leaves were not overtopped by grass leaves. Indeed, in both swards, clover had a greater proportion of its leaf lamina area in the upper, well lit, layers of the canopy than grass did. Consequently, clover had a greater mean rate of leaf photosynthesis in situ in the sward than grass. Clover's advantage in photosynthetic rate per leaf area was offset to some extent by its smaller ratio of leaf area to total above-ground dry weight than grass. The consequences of these results for our understanding of competition between grass and clover in mixed swards are discussed.